Multiple night‐time light‐emitting diode lighting strategies impact grassland invertebrate assemblages

White light‐emitting diodes (LEDs) are rapidly replacing conventional outdoor lighting technologies around the world. Despite rising concerns over their impact on the environment and human health, the flexibility of LEDs has been advocated as a means of mitigating the ecological impacts of globally widespread outdoor night‐time lighting through spectral manipulation, dimming and switching lights off during periods of low demand. We conducted a three‐year field experiment in which each of these lighting strategies was simulated in a previously artificial light naïve grassland ecosystem. White LEDs both increased the total abundance and changed the assemblage composition of adult spiders and beetles. Dimming LEDs by 50% or manipulating their spectra to reduce ecologically damaging wavelengths partially reduced the number of commoner species affected from seven to four. A combination of dimming by 50% and switching lights off between midnight and 04:00 am showed the most promise for reducing the ecological costs of LEDs, but the abundances of two otherwise common species were still affected. The environmental consequences of using alternative lighting technologies are increasingly well established. These results suggest that while management strategies using LEDs can be an effective means of reducing the number of taxa affected, averting the ecological impacts of night‐time lighting may ultimately require avoiding its use altogether.     

The dark side of street lighting: impacts on moths and evidence for the disruption of nocturnal pollen transport

Among drivers of environmental change, artificial light at night is relatively poorly understood, yet is increasing on a global scale. The community‐level effects of existing street lights on moths and their biotic interactions have not previously been studied. Using a combination of sampling methods at matched‐pairs of lit and unlit sites, we found significant effects of street lighting: moth abundance at ground level was halved at lit sites, species richness was >25% lower, and flight activity at the level of the light was 70% greater. Furthermore, we found that 23% of moths carried pollen of at least 28 plant species and that there was a consequent overall reduction in pollen transport at lit sites. These findings support the disruptive impact of lights on moth activity, which is one proposed mechanism driving moth declines, and suggest that street lighting potentially impacts upon pollination by nocturnal invertebrates. We highlight the importance of considering both direct and cascading impacts of artificial light.     

The effects of light pollution on migratory animal behavior

Light pollution is a global threat to biodiversity, especially migratory organisms, some of which traverse hemispheric scales. Research on light pollution has grown significantly over the past decades, but our review of migratory organisms demonstrates gaps in our understanding, particularly beyond migratory birds. Research across spatial scales reveals the multifaceted effects of artificial light on migratory species, ranging from local and regional to macroscale impacts. These threats extend beyond species that are active at night – broadening the scope of this threat. Emerging tools for measuring light pollution and its impacts, as well as ecological forecasting techniques, present new pathways for conservation, including transdisciplinary approaches.     

Conserving energy at a cost to biodiversity? Impacts of LED lighting on bats

Artificial lighting is a key biodiversity threat and produces 1900 million tonnes of CO ₂ emissions globally, more than three times that produced by aviation. The need to meet climate change targets has led to a global increase in energy‐efficient light sources such as high‐brightness light‐emitting diodes (LEDs). Despite the energetic benefits of LEDs, their ecological impacts have not been tested. Using an experimental approach, we show that LED street lights caused a reduction in activity of slow‐flying bats ( Rhinolophus hipposideros and Myotis spp.). Both R. hipposideros and Myotis spp. activities were significantly reduced even during low light levels of 3.6 lux. There was no effect of LED lighting on the relatively fast‐flying Pipistrellus pipistrellus, Pipistrellus pygmaeus and Nyctalus/Eptesicus spp. We provide the first evidence of the effects of LED lights on bats. Despite having considerable energy‐saving benefits, LED lights can potentially fragment commuting routes for bats with associated negative conservation consequences. Our results add to the growing evidence of negative impacts of lighting on a wide range of taxa. We highlight the complexities involved in simultaneously meeting targets for reduction of greenhouse gas emissions and biodiversity loss. New lighting strategies should integrate climate change targets with the cultural, social and ecological impacts of emerging lighting technologies.     

Restless roosts: Light pollution affects behavior,sleep, and physiology in a free‐living songbird

The natural nighttime environment is increasingly polluted by artificial light. Several studies have linked artificial light at night to negative impacts on human health. In free‐living animals, light pollution is associated with changes in circadian, reproductive, and social behavior, but whether these animals also suffer from physiologic costs remains unknown. To fill this gap, we made use of a unique network of field sites which are either completely unlit (control), or are artificially illuminated with white, green, or red light. We monitored nighttime activity of adult great tits, Parus major, and related this activity to within‐individual changes in physiologic indices. Because altered nighttime activity as a result of light pollution may affect health and well‐being, we measured oxalic acid concentrations as a biomarker for sleep restriction, acute phase protein concentrations and malaria infection as indices of immune function, and telomere lengths as an overall measure of metabolic costs. Compared to other treatments, individuals roosting in the white light were much more active at night. In these individuals, oxalic acid decreased over the course of the study. We also found that individuals roosting in the white light treatment had a higher probability of malaria infection. Our results indicate that white light at night increases nighttime activity levels and sleep debt and affects disease dynamics in a free‐living songbird. Our study offers the first evidence of detrimental effects of light pollution on the health of free‐ranging wild animals.     

Artificial night lighting disrupts sex pheromone in a noctuid moth

1. One major, yet poorly studied, change in the environment is the increase in nocturnal light pollution. Although this strongly alters the habitat of nocturnal species, the ecological consequences are poorly known. Moths are well known to be attracted to artificial light sources, but artificial light may affect them in other ways as well. 2. In this study, female Mamestra brassicae moths were subjected to various types of low‐intensity artificial night lighting with contrasting spectral compositions (green‐rich, red‐rich, warm white) or to a dark control treatment and the effects on their sex pheromone production and composition were tested. 3. Artificial night lighting reduced sex pheromone production and altered the chemical composition of the pheromone blend, irrespective of spectral composition. Specifically, amounts of the main pheromone component Z11‐16:Ac were reduced, while the deterring compounds Z9‐14:Ac, Z9‐16:Ac, and Z11‐16:OH were increased relative to Z11‐16:Ac when females were kept under artificial light. These changes may reduce the effectiveness of the sex pheromones, becoming less attractive for males. 4. These results show for the first time that artificial light at night affects processes that are involved in moth reproduction. The potential for mitigation through manipulation of the spectral composition of artificial light appears limited.     

Nest height is affected by lamppost lighting proximity in addition to nestbox size in urban great tits

Both natural and artificial light have proximate influences on many aspects of avian biology, physiology and behaviour. To date artificial light at night is mostly considered as being a nuisance disrupting for instance sleep and reproduction of diurnal species. Here, we investigate if lamppost night lighting affects cavity‐nesting bird species inside their breeding cavity. Nest height in secondary cavity‐nesting species is the result of trade‐offs between several selective forces. Predation is the prevailing force leading birds to build thin nests to increase the distance towards the entrance hole. A thin nest may also limit artificial light exposure at night. Yet, a minimum level of daylight inside nesting cavities is necessary for adequate visual communication and/or offspring development. Against this background, we hypothesised that avian nest‐building behaviour varies in response to a change in night lighting. We monitored nest height of urban great tits Parus major during six years and found that it varied with artificial light proximity. The birds built thinner nests inside nestboxes of various sizes in response to increasing lamppost night light availability at the nest. In large nestboxes, the nests were also thinner when a lamppost was present in the territory. Whether this relationship between artificial night lighting and nest height reflects a positive or negative effect of urbanisation is discussed in the light of recent experimental studies conducted in rural populations by other research groups.     

The ecological impacts of nighttime light pollution: a mechanistic appraisal

The ecological impacts of nighttime light pollution have been a longstanding source of concern, accentuated by realized and projected growth in electrical lighting. As human communities and lighting technologies develop, artificial light increasingly modifies natural light regimes by encroaching on dark refuges in space, in time, and across wavelengths. A wide variety of ecological implications of artificial light have been identified. However, the primary research to date is largely focused on the disruptive influence of nighttime light on higher vertebrates, and while comprehensive reviews have been compiled along taxonomic lines and within specific research domains, the subject is in need of synthesis within a common mechanistic framework. Here we propose such a framework that focuses on the cross‐factoring of the ways in which artificial lighting alters natural light regimes (spatially, temporally, and spectrally), and the ways in which light influences biological systems, particularly the distinction between light as a resource and light as an information source. We review the evidence for each of the combinations of this cross‐factoring. As artificial lighting alters natural patterns of light in space, time and across wavelengths, natural patterns of resource use and information flows may be disrupted, with downstream effects to the structure and function of ecosystems. This review highlights: (i) the potential influence of nighttime lighting at all levels of biological organisation (from cell to ecosystem); (ii) the significant impact that even low levels of nighttime light pollution can have; and (iii) the existence of major research gaps, particularly in terms of the impacts of light at population and ecosystem levels, identification of intensity thresholds, and the spatial extent of impacts in the vicinity of artificial lights.     

Effects of experimental night lighting on the daily timing of winter foraging in common European songbirds

The ecological effects of light pollution are becoming better understood, especially in birds. Recent studies have shown that several bird species can use street lighting to extend activity into the night during the breeding season. However, most of these studies are correlational and little is known about the effects of artificial night lighting on the timing of activities outside the breeding season. During winter, low temperatures and short days may limit foraging opportunities and can negatively affect survival of resident birds. However, night lighting may allow them to expand the time niche available for foraging. Here, we report on a study where we repeatedly manipulated the amount of night lighting during early winter at automated feeding stations in a natural forest. We used video‐recordings at the feeders to determine the time of the first (at dawn) and last (at dusk) foraging visits for six songbird species. We predicted that all species, and in particular the naturally early‐foraging species, would advance their daily onset of foraging during the mornings with night lighting, but would show minimal or no delays in their daily cessation of foraging during the lighted evenings. We found that two early‐foraging species, the blue tit and the great tit, started foraging earlier during the experimentally lighted mornings. However, in great tits, this effect was weak and restricted to nights with inclement weather. The light treatment did not have any effect on the start of foraging in the willow/marsh tit, the nuthatch, the European jay, and the blackbird. Artificial night lighting did not cause later foraging at dusk in any of the six species. Overall, our results suggest that artificial light during winter has only small effects on timing of foraging. We discuss these findings and the importance of temperature and winter weather in shaping the observed foraging patterns.     

Is part‐night lighting an effective measure to limit the impacts of artificial lighting on bats?

As light pollution is currently considered to be a major threat to biodiversity, different lighting management options are being explored to mitigate the impact of artificial lighting on wildlife. Although part‐night lighting schemes have been adopted by many local authorities across Europe to reduce the carbon footprint and save energy, their effects on biodiversity are unknown. Through a paired, in situ experiment, we compared the activity levels of 8 bat species under unlit, part‐night, and full‐night lighting treatments in a rural area located 60 km south of Paris, France. We selected 36 study locations composed of 1 lit site and a paired unlit control site; 24 of these sites were located in areas subject to part‐night lighting schemes, and 12 sites were in areas under standard, full‐night lighting. There was significantly more activity on part‐night lighting sites compared to full‐night lighting sites for the late‐emerging, light‐sensitive Plecotus spp., and a similar pattern was observable for Myotis spp., although not significant. In contrast, part‐night lighting did not influence the activity of early emerging bat species around streetlights, except for Pipistrellus pipistrellus for which there was significantly less activity on part‐night lighting sites than on full‐night lighting sites. Overall, no significant difference in activity between part‐ and full‐night lighting sites were observed in 5 of the 8 species studied, suggesting that current part‐night lighting schemes fail to encompass the range of activity of most bat species. We recommend that such schemes start earlier at night to effectively mitigate the adverse effects of artificial lighting on light‐sensitive species, particularly along ecological corridors that are especially important to the persistence of biodiversity in urban landscapes.     

Invasive house geckos are more willing to use artificial lights than are native geckos

There is increasing concern about the ecological effects of light pollution, as artificial lighting spreads with urban expansion. While artificial lighting can negatively affect some species, others use it in novel ways. In tropical and subtropical regions, artificial lighting has created a novel niche: the ‘night light’ niche. Geckos living as human commensals (house geckos) are apparently well adapted to occupy this niche. In an urban area in north‐eastern Australia, we found that the invasive Asian house gecko Hemidactylus frenatus (Gekkonidae) occupies a broader range of light environments in the field than does the native gecko Gehyra dubia (Gekkonidae). Experimental removal of the invasive species from a building indicated that it did not behaviourally influence the light environments chosen by the native species in the short term; they continued to use darker areas even after the invasive species was removed. In Y‐maze experiments, neither species showed a significant preference for light or dark areas; however, preliminary data suggest the invasive species was more willing to explore the Y‐maze than the native species. The willingness of H. frenatus to forage closer to lights, where insect abundance is typically higher, might account for its success as a global invader of human environments, even in areas where other gecko species are established.     

Declines in moth populations stress the need for conserving dark nights

Given the global continuous rise, artificial light at night is often considered a driving force behind moth population declines. Although negative effects on individuals have been shown, there is no evidence for effects on population sizes to date. Therefore, we compared population trends of Dutch macromoth fauna over the period 1985–2015 between moth species that differ in phototaxis and adult circadian rhythm. We found that moth species that show positive phototaxis or are nocturnally active have stronger negative population trends than species that are not attracted to light or are diurnal species. Our results indicate that artificial light at night is an important factor in explaining declines in moth populations in regions with high artificial night sky brightness. Our study supports efforts to reduce the impacts of artificial light at night by promoting lamps that do not attract insects and reduce overall levels of illumination in rural areas to reverse declines of moth populations.     

Lamp-Lit Bridges as Dual Light-Traps for the Night-Swarming Mayfly,Ephoron virgo: Interaction of Polarized and Unpolarized Light Pollution

Ecological photopollution created by artificial night lighting can alter animal behavior and lead to population declines and biodiversity loss. Polarized light pollution is a second type of photopollution that triggers water-seeking insects to ovisposit on smooth and dark man-made objects, because they simulate the polarization signatures of natural water bodies. We document a case study of the interaction of these two forms of photopollution by conducting observations and experiments near a lamp-lit bridge over the river Danube that attracts mass swarms of the mayfly Ephoron virgo away from the river to oviposit on the asphalt road of the bridge. Millions of mayflies swarmed near bridge-lights for two weeks. We found these swarms to be composed of 99% adult females performing their upstream compensatory flight and were attracted upward toward unpolarized bridge-lamp light, and away from the horizontally polarized light trail of the river. Imaging polarimetry confirmed that the asphalt surface of the bridge was strongly and horizontally polarized, providing a supernormal ovipositional cue to Ephoron virgo, while other parts of the bridge were poor polarizers of lamplight. Collectively, we confirm that Ephoron virgo is independently attracted to both unpolarized and polarized light sources, that both types of photopollution are being produced at the bridge, and that spatial patterns of swarming and oviposition are consistent with evolved behaviors being triggered maladaptively by these two types of light pollution. We suggest solutions to bridge and lighting design that should prevent or mitigate the impacts of such scenarios in the future. The detrimental impacts of such scenarios may extend beyond Ephoron virgo.     

Pollination by nocturnal Lepidoptera,and the effects of light pollution: a review

1. Moths (Lepidoptera) are the major nocturnal pollinators of flowers. However, their importance and contribution to the provision of pollination ecosystem services may have been under‐appreciated. Evidence was identified that moths are important pollinators of a diverse range of plant species in diverse ecosystems across the world. 2. Moth populations are known to be undergoing significant declines in several European countries. Among the potential drivers of this decline is increasing light pollution. The known and possible effects of artificial night lighting upon moths were reviewed, and suggest how artificial night lighting might in turn affect the provision of pollination by moths. The need for studies of the effects of artificial night lighting upon whole communities of moths was highlighted. 3. An ecological network approach is one valuable method to consider the effects of artificial night lighting upon the provision of pollination by moths, as it provides useful insights into ecosystem functioning and stability, and may help elucidate the indirect effects of artificial light upon communities of moths and the plants they pollinate. 4. It was concluded that nocturnal pollination is an ecosystem process that may potentially be disrupted by increasing light pollution, although the nature of this disruption remains to be tested.     

Nocturnal city lighting elicits a macroscale response from an insect outbreak population

Anthropogenic environmental change affects organisms by exposing them to enhanced sensory stimuli that can elicit novel behavioural responses. A pervasive feature of the built environment is artificial nocturnal lighting, and brightly lit urban areas can influence organism abundance, distribution and community structure within proximate landscapes. In some cases, the attractive or disorienting effect of artificial light at night can draw animals into highly unfavourable habitats, acting as a macroscale attractive ecological sink. Despite their significance for animal ecology, identifying cases of these phenomena and determining their effective scales and the number of organisms impacted remains challenging. Using an integrated set of remote-sensing observations, we quantify the effect of a large-scale attractive sink on nocturnal flights of an outbreak insect population in Las Vegas, USA. At the peak of the outbreak, over 45 million grasshoppers took flight across the region, with the greatest numbers concentrating over high-intensity city lighting. Patterns of dusk ascent from vegetated habitat toward urban areas suggest a daily pull toward a time-varying nocturnal attractive sink. The strength of this attractor varies with grasshopper density. These observations provide the first macroscale characterization of the effects of nocturnal urban lighting on the behaviour of regional insect populations and demonstrate the link between insect perception of the built environment and resulting changes in spatial and movement ecology. As human-induced environmental change continues to affect insect populations, understanding the impacts of nocturnal light on insect behaviour and fitness will be vital to developing robust large-scale management and conservation strategies.     

Guiding lights: Foraging responses of juvenile nocturnal orb‐web spiders to the presence of artificial light at night

The reach of artificial light at night (ALAN) is growing rapidly around the globe, including the increasing use of energy‐efficient LED lights. Many studies document the physiological costs of light at night, but far fewer have focused on the potential benefits for nocturnal insectivores and the likely ecological consequences of shifts in predator–prey relationships. We investigated the effects of ALAN on the foraging behaviour and prey capture success in juvenile Australian garden orb‐web spiders (Eriophora biapicata). Laboratory experiments demonstrated that juvenile spiders were attracted to LED lights when choosing foraging sites, but prey availability was a stronger cue for remaining in a foraging site. Field experiments revealed a significant increase in prey capture rates for webs placed near LED lights. This suggests that any physiological costs of light at night may be offset by the foraging benefits, perhaps partially explaining recently observed increases in the size, fecundity and abundance of some orb‐web spider species in urban environments. Our results highlight the potential long‐term consequences of night lighting in urban ecosystems, through the impact of orb‐web spiders on insect populations.     

Systematic review of light exposure impact on human circadian rhythm

Light is necessary for life, and artificial light improves visual performance and safety, but there is an increasing concern of the potential health and environmental impacts of light. Findings from a number of studies suggest that mistimed light exposure disrupts the circadian rhythm in humans, potentially causing further health impacts. However, a variety of methods has been applied in individual experimental studies of light-induced circadian impacts, including definition of light exposure and outcomes. Thus, a systematic review is needed to synthesize the results. In addition, a review of the scientific evidence on the impacts of light on circadian rhythm is needed for developing an evaluation method of light pollution, i.e., the negative impacts of artificial light, in life cycle assessment (LCA). The current LCA practice does not have a method to evaluate the light pollution, neither in terms of human health nor the ecological impacts. The systematic literature survey was conducted by searching for two concepts: light and circadian rhythm. The circadian rhythm was searched with additional terms of melatonin and rapid-eye-movement (REM) sleep. The literature search resulted to 128 articles which were subjected to a data collection and analysis. Melatonin secretion was studied in 122 articles and REM sleep in 13 articles. The reports on melatonin secretion were divided into studies with specific light exposure (101 reports), usually in a controlled laboratory environment, and studies of prevailing light conditions typical at home or work environments (21 studies). Studies were generally conducted on adults in their twenties or thirties, but only very few studies experimented on children and elderly adults. Surprisingly many studies were conducted with a small sample size: 39 out of 128 studies were conducted with 10 or less subjects. The quality criteria of studies for more profound synthesis were a minimum sample size of 20 subjects and providing details of the light exposure (spectrum or wavelength; illuminance, irradiance or photon density). This resulted to 13 qualified studies on melatonin and 2 studies on REM sleep. Further analysis of these 15 reports indicated that a two-hour exposure to blue light (460 nm) in the evening suppresses melatonin, the maximum melatonin-suppressing effect being achieved at the shortest wavelengths (424 nm, violet). The melatonin concentration recovered rather rapidly, within 15 min from cessation of the exposure, suggesting a short-term or simultaneous impact of light exposure on the melatonin secretion. Melatonin secretion and suppression were reduced with age, but the light-induced circadian phase advance was not impaired with age. Light exposure in the evening, at night and in the morning affected the circadian phase of melatonin levels. In addition, even the longest wavelengths (631 nm, red) and intermittent light exposures induced circadian resetting responses, and exposure to low light levels (5–10 lux) at night when sleeping with eyes closed induced a circadian response. The review enables further development of an evaluation method of light pollution in LCA regarding the light-induced impacts on human circadian system.     

Environmental and human health impacts of growing genetically modified herbicide-tolerant sugar beet: a life-cycle assessment

There is ongoing debate concerning the possible environmental and human health impacts of growing genetically modified (GM) crops. Here, we report the results of a life-cycle assessment (LCA) comparing the environmental and human health impacts of conventional sugar beet growing regimes in the UK and Germany with those that might be expected if GM herbicide-tolerant (to glyphosate) sugar beet is commercialized. The results presented for a number of environmental and human health impact categories suggest that growing the GM herbicide-tolerant crop would be less harmful to the environment and human health than growing the conventional crop, largely due to lower emissions from herbicide manufacture, transport and field operations. Emissions contributing to negative environmental impacts, such as global warming, ozone depletion, ecotoxicity of water and acidification and nutrification of soil and water, were much lower for the herbicide-tolerant crop than for the conventional crop. Emissions contributing to summer smog, toxic particulate matter and carcinogenicity, which have negative human health impacts, were also substantially lower for the herbicide-tolerant crop. The environmental and human health impacts of growing GM crops need to be assessed on a case-by-case basis using a holistic approach. LCA is a valuable technique for helping to undertake such assessments.     

Artificial light at night causes diapause inhibition and sex‐specific life history changes in a moth

Rapidly increasing levels of light pollution subject nocturnal organisms to major alterations of their habitat, the ecological consequences of which are largely unknown. Moths are well‐known to be attracted to light at night, but effects of light on other aspects of moth ecology, such as larval development and life‐history, remain unknown. Such effects may have important consequences for fitness and thus for moth population sizes. To study the effects of artificial night lighting on development and life‐history of moths, we experimentally subjected Mamestra brassicae (Noctuidae) caterpillars to low intensity green, white, red or no artificial light at night and determined their growth rate, maximum caterpillar mass, age at pupation, pupal mass and pupation duration. We found sex‐specific effects of artificial light on caterpillar life‐history, with male caterpillars subjected to green and white light reaching a lower maximum mass, pupating earlier and obtaining a lower pupal mass than male caterpillars under red light or in darkness. These effects can have major implications for fitness, but were absent in female caterpillars. Moreover, by the time that the first adult moth from the dark control treatment emerged from its pupa (after 110 days), about 85% of the moths that were under green light and 83% of the moths that were under white light had already emerged. These differences in pupation duration occurred in both sexes and were highly significant, and likely result from diapause inhibition by artificial night lighting. We conclude that low levels of nocturnal illumination can disrupt life‐histories in moths and inhibit the initiation of pupal diapause. This may result in reduced fitness and increased mortality. The application of red light, instead of white or green light, might be an appropriate measure to mitigate negative artificial light effects on moth life history.     

The intensity and spectrum of artificial light at night alters crayfish interactions

ABSTRACT

Ecological light pollution (ELP) is quickly becoming a worldwide concern and can negatively affect aquatic ecosystems. The given intensity and spectrum of a light source can influence how organisms function within their environment. These properties of artificial lighting at night (ALAN) and their impacts on the physiology and behaviour of crayfish were examined in this work. Hemolymph was obtained from crayfish to quantify a physiological response. Behavioural data were measured as the number, duration, and maximum intensity of agonistic fights. Exposure to higher intensities of light and the presence of ultraviolet light induced a behavioural trend, resulting in significantly altered social interactions within both species of crayfish. The number and maximum intensity of lights significantly decreased, whereas the duration of time spent fighting significantly increased. Due to the importance of freshwater environments and the role crayfish play as a keystone species, examining how crayfish are impacted from ALAN is imperative to maintaining the health of aquatic ecosystems.